C3H. altered development COMT. N. tabacum Antisense 42 Reduced b No changes e Larger xylem cells Reduced a S/G increased f

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1 TABLE 1 Overexpression, down-regulation, and mutation of monolignol biosynthesis genes. Gene Plant Type of Enzyme Lignin Lignin Plant development Pathogen Stem References modification activity in content composition sensitivity color stems (%) PAL Nicotiana tabacum Sense 2.3 Reduced a S/G increased g Altered development bis S and G decreased g Sense 10 Reduced a,b S/G increased f Altered development, increased Increased susceptibility - 3, 17, 18, 40, 54,55 G decreased f systemic resistance to insect to Cercospora nicotianae larvae (Heliothis virescens) fungal infection, SAR inhibited after TMV infection Sense 150 Increased a S/G decreased g bis S decreased g, Sense 200 No changes a No changes f Loss of systemic resistance SAR enhanced after TMV - 18, 28, 54,55 to insect larvae (H. virescens infection or Manduca sexta) C4HI N. tabacum Antisense 20 Reduced a S/G decreased f No changes S and G decreased f Sense 200 No changes a No changes f No changes C4HII N. tabacum Antisense and sense 10 Reduced d S/G decreased g No changes S and G decreased g C3H COMT Arabidopsis thaliana Mutant (ref8) - Reduced a,b Trace of S and G units e,f,k Collapsed vessels, Increased fungal colonization - 19, 20 more H units e,f,k altered development N. tabacum Antisense 42 Reduced b No changes e Larger xylem cells Reduced a S/G increased f S and G decreased f Antisense 54 No changes c Sy/V decreased e No changes Sy decreased e Sense 2 No changes a S decreased g, 5OHG g 2 S/G decreased g No changes g Populus tremula x P. alba Antisense 5 No changes a S/G decreased g No changes No increased pale rose 37, 45, 50, 51, 58 S decreased g, damage caused by G increased g, 5OHG g feeding insects in

2 CCoAOMT the field Sense <3 Reduced a S/G decreased g, 5OHG g, No changes - brown 31, 50, 51 S decreased g, more coniferaldehyde g P. tremuloides Sense 28 No changes a,d S/G decreased g, 5OHG g, No changes - red-brown 57 G increased g, S decreased g more coniferaldehyde g Medicago sativa Antisense 4.5 Reduced a, G decreased g, no S g, No changes - 22 No changes d S/G decreased g Sense 15 Reduced a,, S and G decreased g, No changes d S/G decreased g, 5OHG g Zea mays Mutant (bm3) 10 Reduced a S/G decreased g, 5OHG g, No changes - brown-red 10, 10bis, 11, 59, S g and Syr e decreased 21bis, 33ter, 36bis Antisense 15 Reduced a S decreased g, 5OHG g No changes - brown-red 48bis G increased g Sorghum bicolor Mutant (bmr12 and - Reduced a G increased g, S decreased g, - - brown-red 9bis bmr18) 5OHG g, S/G decreased g N. tabacum Antisense 25 Reduced a Sy/V increased e Altered vessel shape Sy and V decreased e Antisense ~20 Reduced a No changes g Reduced growth altered development P. tremula x P.alba Sense 10% protein Reduced a S/G increased g No changes - pink-red 43 Amount S and G decreased g, more p-hydroxybenzoic acid h Sense 30 Reduced a,m S and G decreased f Altered vessel shape - light orange 62 M. sativa Antisense 4 Reduced a,d G decreased g, No changes S/G increased g CCoAOMT/COMT N. tabacum Antisense/antisense 8/14 Reduced a S/G decreased f Altered vessel shape S and G decreased f Antisense/antisense ~20/~30 Reduced a S decreased g, 5OHG g Reduced growth, altered Larger necrotic lesions - 27,47 development upon TMV infection M. sativa Antisense/antisense 5/12 Reduced a S decreased g, No changes No changes d S/G decreased g Sense/sense 25/11 Reduced a S and G decreased g, No changes No changes d S/G decreased g F5H N. tabacum Sense - Reduced a,b Sy/V increased e, No changes or lignin more Sy e or S increased k, extractable V decreased e

3 A. thaliana Sense - Reduced a Sy e or S, h,k increased No changes , 42 V e or G k decreased Mutant (fah1) - No changes a Trace levels of Syr e, No changes , 41, 42, 56bis Sy e and S h,k, Van e, V e or G increased k Sense - Reduced a S increased g, G decreased g No changes bis P. tremula x P. alba Sense - - S/G increased k No changes Sy e decreased or S k increased, V decreased e 4CL N. tabacum Sense <1 Reduced a,d Syr/Van decreased or No changes - brown 32, 33 or increased e Van and Syr decreased e, p-hydroxybenzaldehyde increased n, H increased n, S and G decreased n more HCAs h,i Antisense ~20 Reduced d Sy/V increased e, No changes - brown 33 less aldehydes e A. thaliana Antisense 8 Reduced b V decreased e No changes Sy/V increased e P. tremuloides Antisense 10 Reduced a No changes g Increased growth CCR N. tabacum Antisense 30 Reduced a,d S and G decreased g, Collapsed vessels, - orange brown 7, 8, 48,49 S/G increased g, reduced growth tyramine ferulate increased h lower vessel cell area Sense 1 Reduced a S and G decreased g abnormal leaf, - orange brown 44bis S/G increased g reduced growth P. tremula x P. alba Sense and antisense - Reduced a S/G increased g Collapsed vessels - orange J.-C. Leplé, C. Lapierre and W. Boerjan A. thaliana Mutant (irx4) - Reduced b,h - Collapsed vessels, reduced growth COMT/CCR N. tabacum Antisense/antisense 41/10 Reduced a S/G increased g Reduced growth CAD2 N. tabacum Antisense 7 No changes a,b,d More aldehydes f Reduced vessel cell area - red brown 7, 8, 24 S/G decreased g Antisense 8 No changes a,d S/G decreased g, No changes - red 49, 60 more cinnamaldehydes h Antisense 9 No changes d More aldehydes j No changes - red 56

4 Antisense 45 No changes d More aldehydes g No changes - brown 25, 26 P. tremula x P. alba Antisense 30 Reduced a More Syr g No changes No increased red 5, 37, 45 damage caused by feeding insects in the field Sense 30 No changes a No changes g No changes - red 5, 37 M. sativa Antisense 30 No changes a More aldehydes, No changes - red 4 S/G decreased g S decreased g Z. mays Mutant (bm1) ~30 Reduced a Lignin more condensed g No changes - brown-red 23 S and G decreased g S. bicolor Mutant (bmr6) ~30 Reduced a,b S and G decreased f,g, No changes - brown-red 9bis, 45bis coniferaldehyde increased f,g Van and Syr decreased e Pinus taeda Mutant (cad-n1) 1 Reduced a More dihydroconiferyl No changes - brown 39, 52 alcohol h, more coniferaldehyde h CAD2/COMT N. tabacum Antisense/antisense Variable/variable No changes a,d No changes g No changes - No changes 1 Sense/sense 15/36 Reduced a,d - No changes - bronze red 1 CAD2/CCR N. tabacum Antisense/antisense 12/32 Reduced a S/G increased g Reduced vessel cell area - brown 7, 8 S and G decreasedg tyramine ferulate increased h CAD2/COMT/CCR N. tabacum Sense/Sense/Sense 4/24/18 Reduced a,d - Collapsed vessels - bronze red 1 Altered growth POX N. tabacum Sense 250 Increased b,d No changes g Wilting Greater resistance - 9, 34, 36 Altered development to insects Antisense 1 No changes b,d No changes g Taller growth, early flowering - - 9, 35 Sense increased No changes a More coniferaldehyde g More flexible stems Increased susceptibility No changes 16 to Phytophthora parasitica var. nicotianae, suppression of Erwinia carotovora growth P. tremula x P. alba Sense 800 No changes a No changes g No changes - No changes 12 P. kitakamiensis Antisense 56 Reduced d,l S/G increased g,l No changes - No changes 61 Lycopersicon esculentum Sense - Increased b - Altered development - No changes 15 LAC P. tremula x P. alba Antisense - No changes a,d No changes g Xylem fiber wall alterations Liriodendron tulipifera Antisense - No changes No changes

5 a Klason. b Thioglycolic acid. c Sum of Klason and acid-soluble lignin content. d Acetyl bromide. e Nitrobenzene oxidation. f Pyrolysis gas chromatography-mass spectrometry. g Thioacidolysis. h NMR. i Alkaline hydrolysis followed by gas chromatography. j Fourier transform infra red. k Derivatization followed by reductive cleavage. l Permanganate oxidation. m Diffuse reflectance infrared Fourier transform spectroscopy n Pyrolysis gas chromatography Antisense, transgenic plants with an antisense construct; G, guaiacyl; H, p-hydroxyphenyl; HCA, hydroxycinnamic acid; 5OHG, 5-hydroxyguaiacyl; LAC, laccase; POX, peroxidase; S, syringyl; SAR, systemic acquired resistance; sense, transgenic plants with a sense construct; Sy, sum of Syr and syringic acid; Syr, syringaldehyde; TMV, tobacco mosaic virus; V, sum of Van and vanillic acid; Van, vanillin; -, not determined. The data on lignin composition reported in this Table are restricted to S/G ratio and the incorporation of unusual phenolics.

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8 29. Hu W-J, Harding SA, Lung J, Popko JL, Ralph J, et al Repression of lignin biosynthesis promotes cellulose accumulation and growth in transgenic trees. Nature Biotechnol. 17: Jones L, Ennos AR, Turner SR Cloning and characterization of irregular xylem4 (irx4): a severely lignin-deficient mutant of Arabidopsis. Plant J. 26: Jouanin L, Goujon T, de Nadaï V, Martin M-T, Mila I, et al Lignification in transgenic poplars with extremely reduced caffeic acid O-methyltransferase activity. Plant Physiol. 123: Kajita S, Hishiyama S, Tomimura Y, Katayama Y, Omori S Structural characterization of modified lignin in transgenic tobacco plants in which the activity of 4-coumarate:coenzyme A ligase is depressed. Plant Physiol. 114: Kajita S, Katayama Y, Omori S Alterations in the biosynthesis of lignin in transgenic plants with chimeric genes for 4-coumarate:coenzyme A ligase. Plant Cell Physiol. 37: bis 33ter Korth KL, Blount JW, Chen F, Rasmussen S, Lamb C, Dixon RA Changes in phenylpropanoid metabolites associated with homology-dependent silencing of phenylalanine ammonia-lyase and its somatic reversion in tobacco. Physiol. Plant. 111: Kuć J, Nelson OE, Flanagan P Degradation of abnormal lignins in the brown-midrib mutants and double mutants of maize. Phytochemistry 7: Lagrimini LM Wound-induced deposition of polyphenols in transgenic plants overexpressing peroxidase. Plant Physiol. 96: Lagrimini LM, Gingas V, Finger F, Rothstein S, Liu T-TY Characterization of antisense transformed plants deficient in the tobacco anionic peroxidase. Plant Physiol. 114: Lagrimini LM, Joly RJ, Dunlap JR, Liu T-TY The consequence of peroxidase overexpression in transgenic plants on root growth and development. Plant Mol. Biol. 33: bis Lapierre C, Tollier M-T, Monties, B Mise en évidence d'un nouveau type d'unité constitutive dans les lignines d'un mutant de Maïs bm3. C.R. Acad. Sci. Paris, Série III, 307: Lapierre C, Pollet B, Petit-Conil M, Toval G, Romero J, et al Structural alterations of lignins in transgenic poplars with depressed cinnamyl alcohol dehydrogenase or caffeic acid O-methyltransferase activity have opposite impact on the efficiency of industrial Kraft pulping. Plant Physiol. 119: Lee D, Meyer K, Chapple C, Douglas CJ Antisense suppression of 4-coumarate:coenzyme A ligase activity in Arabidopsis leads to altered lignin subunit composition. Plant Cell 9: MacKay JJ, O'Malley DM, Presnell T, Booker FL, Campbell MM, Whetten RW, Sederoff RR Inheritance, gene expression, and lignin characterization in a mutant pine deficient in cinnamyl alcohol dehydrogenase. Proc. Natl. Acad. Sci. USA 94: Maher EA, Bate NJ, Ni W, Elkind Y, Dixon RA, Lamb CJ Increased disease susceptibility of transgenic tobacco plants with suppressed levels of preformed phenylpropanoid products. Proc. Natl. Acad. Sci. USA 91: Marita JM, Ralph J, Hatfield RD, Chapple C NMR characterization of lignins in Arabidopsis altered in the activity of ferulate 5-hydroxylase. Proc. Natl. Acad. Sci. USA 96: Meyer K, Shirley AM, Cusumano JC, Bell-Lelong DA, Chapple C Lignin monomer composition is determined

9 by the expression of a cytochrome P450-dependent monooxygenase in Arabidopsis. Proc. Natl. Acad. Sci. USA 95: Meyermans H, Morreel K, Lapierre C, Pollet B, De Bruyn A, et al Modification in lignin and accumulation of phenolic glucosides in poplar xylem upon down-regulation of caffeoyl-coenzyme A O-methyltransferase, an enzyme involved in lignin biosynthesis. J. Biol. Chem. 275: Ni W, Paiva NL, Dixon RA Reduced lignin in transgenic plants containing a caffeic acid O-methyltransferase antisense gene. Transgenic Res. 3: bis O Connell A, Holt K, Piquemal J, Grima-Pettenati J, Boudet A, et al Improved paper pulp from plants with suppressed cinnamoyl-coa reductase or cinnamyl alcohol dehydrogenase. Transgenic Res. 11: Pilate G, Guiney E, Holt K, Petit-Conil M, Lapierre C, et al Field and pulping performances of transgenic trees with altered lignification. Nature Biotechnol. 20: bis Pillonel C, Mulder MM, Boon JJ, Forster B, Binder A Involvement of cinnamyl-alcohol dehydrogenase in the control of lignin formation in Sorghum bicolor L. Moench. Planta 185: Pinçon G, Chabannes M, Lapierre C, Pollet B, Ruel K, et al Simultaneous down-regulation of caffeic/5-hydroxy ferulic acid-o-methyltransferase I and cinnamoyl-coenzyme A reductase in the progeny from a cross between tobacco lines homozygous for each transgene. Consequences for plant development and lignin synthesis. Plant Physiol. 126: Pinçon G, Maury S, Hoffmann L, Geoffroy P, Lapierre C, Pollet B, Legrand M Repression of O-methyltransferase genes in transgenic tobacco affects lignin synthesis and plant growth. Phytochemistry 57: Piquemal J, Lapierre C, Myton K, O'Connell A, Schuch W, Grima-Pettenati J, Boudet A-M Down-regulation in cinnamoyl-coa reductase induces significant changes of lignin profiles in transgenic tobacco plants. Plant J. 13: bis Piquemal J, Chamayou S, Nadaud I, Beckert M, Barrière Y, Mila I, Lapierre C, Rigau J, Puigdomenech P, Jauneau A, Digonnet C, Boudet A-M, Goffner D, Pichon M Down-regulation of caffeic acid O-methyltransferase in maize revisited using a transgenic approach. Plant Physiol. 130: Ralph J, Hatfield RD, Piquemal J, Yahiaoui N, Pean M, Lapierre C, Boudet AM NMR characterization of altered lignins extracted from tobacco plants down-regulated for lignification enzymes cinnamyl-alcohol dehydrogenase and cinnamoyl-coa reductase. Proc. Natl. Acad. Sci. USA 95: Ralph J, Lapierre C, Lu F, Marita JM, Pilate G, et al NMR evidence for benzodioxane structures resulting from incorporation of 5-hydroxyconiferyl alcohol into lignins of O-methyltransferase-deficient plants. J. Agric. Food Chem. 49: Ralph J, Lapierre C, Marita JM, Kim H, Lu F, et al Elucidation of new structures in lignins of CAD- and COMT-deficient plants by NMR. Phytochemistry 57: Ralph J, MacKay JJ, Hatfield RD, O'Malley DM, Whetten RW, Sederoff RR Abnormal lignin in a loblolly pine mutant. Science 277: Ranocha P, Chabannes M, Chamayou S, Danoun S, Jauneau A, Boudet A-M, Goffner D Laccase down-regulation causes alterations in phenolic metabolism and cell wall structure in poplar. Plant Physiol. 129: Sewalt VJH, Ni W, Blount JW, Jung HG, Masoud SA, et al Reduced lignin content and altered lignin composition in transgenic tobacco down-regulated in expression of L-phenylalanine ammonia-lyase or cinnamate 4-hydroxylase. Plant Physiol. 115:41-50

10 55. Sewalt VJH, Ni W, Jung H, Dixon RA Lignin impact on fiber degradation: increased enzymatic digestibility of genetically engineered tobacco (Nicotina tabacum) stems reduced in lignin content. J. Agric. Food Chem. 45: Stewart D, Yahiaoui N, McDougall GJ, Myton K, Marque C, Boudet AM, Haigh J Fourier-transform infrared and Raman spectroscopic evidence for the incorporation of cinnamaldehydes into the lignin of transgenic tobacco (Nicotiana tabacum L.) plants with reduced expression of cinnamyl alcohol dehydrogenase. Planta 201: bis Sibout R, Baucher M, Van Doorsselaere J, Gatineau M, Pollet B, Mila I, Maba B, Pilate G, Lapierre C, Boerjan W, Jouanin L Expression of a poplar ferulate-5-hydroxylase in Arabidopsis thaliana. Plant Physiol. Biochem. 40: Tsai C-J, Popko JL, Mielke MR, Hu W-J, Podila GK, Chiang V.L Suppression of O-methyltransferase gene by homologous sense transgene in quaking aspen causes red-brown wood phenotypes. Plant Physiol. 117: Van Doorsselaere J, Baucher M, Chognot E, Chabbert B, Tollier M-T, et al A novel lignin in poplar trees with a reduced caffeic acid/5-hydroxyferulic acid O-methyltransferase activity. Plant J. 8: Vignols F, Rigau J, Torres MA, Capellades M, Puigdomènech P The brown midrib3 (bm3) mutation in maize occurs in the gene encoding caffeic acid O-methyltransferase. Plant Cell 7: Yahiaoui N, Marque C, Myton KE, Negrel J, Boudet AM Impact of different levels of cinnamyl alcohol dehydrogenase down-regulation on lignins of transgenic tobacco plants. Planta 204: Yahong L, Tsuji Y, Nishikubo N, Kajita S, Morohoshi N Analysis of transgenic poplar in which the expression of peroxidase gene is suppressed. See ref. 43a. pp Zhong R, Morrison WH III, Himmelsbach DS, Poole FL II, Ye Z-H Essential role of caffeoyl coenzyme A O-methyltransferase in lignin biosynthesis in woody poplar plants. Plant Physiol. 124: Zhong R, Morrison WH III, Negrel J, Ye Z-H Dual methylation pathways in lignin biosynthesis. Plant Cell 10:

Signatures of cinnamyl alcohol dehydrogenase deficiency in poplar lignins

Phytochemistry 65 (2004) 313 321 www.elsevier.com/locate/phytochem Signatures of cinnamyl alcohol dehydrogenase deficiency in poplar lignins Catherine Lapierre a, *, Gilles Pilate b, Brigitte Pollet a,